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geodiversitas
2022 z 44 z 1
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The evolution of hermit crabs (Crustacea, Decapoda,
Anomura, Paguroidea) on the basis of carapace
morphology: a state-of-the-art-report
René H. B. FRAAIJE
Barry W. M. VAN BAKEL
Oertijdmuseum, Bosscheweg 80, 5283 WB Boxtel (the Netherlands)
info@oertijdmuseum.nl (corresponding author)
barryvanbakel@gmail.com
John W. M. JAGT
Natuurhistorisch Museum Maastricht,
De Bosquetplein 6-7, 6211 KJ Maastricht (the Netherlands)
john.jagt@maastricht.nl
Sylvain CHARBONNIER
Centre de Recherche en Paléontologie – Paris (UMR 7207 CR2P),
CNRS-MNHN-Sorbonne Université,
Muséum national d’Histoire naturelle, Département Origines & Évolution,
case postale 38, 57 rue Cuvier, 75231 Paris cedex 05 (France)
sylvain.charbonnier@mnhn.fr
Guenter SCHWEIGERT
Staatliches Museum für Naturkunde, Rosenstein 1, 70191 Stuttgart (Germany)
guenter.schweigert@smns-bw.de
Géraldine GARCIA
PALEVOPRIM, CNRS-UMR 7262 CNRS, INEE, Université de Poitiers,
6 rue Michel Brunet, 86073 Poitiers (France)
geraldine.garcia@univ-poitiers.fr
Xavier VALENTIN
PALEVOPRIM, CNRS-UMR 7262 CNRS, INEE, Université de Poitiers,
6 rue Michel Brunet, 86073 Poitiers (France)
and Palaios, Research Association, 15 rue de l’Aumônerie, 86300 Valdivienne (France)
xavier.valentin@univ-poitiers.fr
Submitted on 9 June 2020 | accepted on 14 October 2020 | published on 6 January 2022
urn:lsid:zoobank.org:pub:6A4424D0-DEFD-4783-9B75-24AC2C7829F7
Fraaije R. H. B., Van Bakel B. W. M., Jagt J. W. M., Charbonnier S., Schweigert G., Garcia G. & Valentin X. 2022. — The
evolution of hermit crabs (Crustacea, Decapoda, Anomura, Paguroidea) on the basis of carapace morphology: a state-of-
the-art-report. Geodiversitas 44 (1): 1-16. https://doi.org/10.5252/geodiversitas2022v44a1. http://geodiversitas.com/44/1
ABSTRACT
In our ongoing studies of both extinct and extant hermit crabs (Paguroidea Latreille, 1802), we have
observed and recorded a number of morphological changes that have taken place between Early Jurassic
times (c. 185 million years ago) and the present day. Species compositions of paguroid assemblages from
GEODIVERSITAS • 2022 • 44 (1) © Publications scientifiques du Muséum national d’Histoire naturelle, Paris. www.geodiversitas.com 1
Fraaije R. H. B. et al.
marine Upper Jurassic and Lower Cretaceous rocks in Europe are here compared with those of modern
marine settings. Basal paguroids with cylindrical carapaces predominated in reefal environments
during the Late Jurassic, but were gradually replaced by hermit crabs with non-cylindrical carapaces.
The most basal paguroids reveal a branchial groove, but branchial condensation towards a single (i.e.,
cervical) groove is here shown to have occurred early in their evolutionary history. In several extant,
genera remnants of a branchial groove, in combination with several other basal features, can be seen.
For this reason, these are here considered to be the most primitive members amongst extant paguroids.
In this study, we resurrect the family Probeebeidae and erect a new family, Paguropsidae n. fam, to
accommodate extant basal hermit crabs such as Paguropsis Henderson, 1888, Eopaguropsis Fraaije,
Krzemiński, Van Bakel, Krzemińska & Jagt, 2012 and Paguropsina Lemaitre, Rahayu & Komai,
2018. Described are also one pair of Early Jurassic (Pliensbachian) and two pairs of Middle Jurassic
KEY WORDS (Callovian) isochelous paguroid chelae, all collected recently. A new specimen of Schobertella reveals
Branchial condensation,
isochely, some important morphological traits of the shield that had not been observed previously. The earliest
heterochely, known example to date of clearly heterochelous chelae in the fossil record of hermit crabs originates from
Probeebeidae,
Paguropsidae n. fam, upper Kimmeridgian/Tithonian strata in the Boulonnais (northwestern France). For the first time, a
new family. phylogenetic scheme of marine Paguroidea, as based on morphological data of carapaces, is presented.
RÉSUMÉ
L’évolution des bernard-l’hermite (Crustacea, Decapoda, Anomura, Paguroidea) à partir de la morpho-
logie des carapaces : état des connaissances.
Dans nos études en cours sur les bernard-l’hermite fossiles et actuels (Paguroidea Latreille, 1802),
nous avons observé et reconnu un certain nombre de changements morphologiques qui ont eu lieu
entre le début du Jurassique (c. 185 millions d’années) et aujourd’hui. Les compositions d’espèces
des assemblages de paguroïdes provenant de roches marines du Jurassique supérieur et du Crétacé
inférieur d’Europe sont ici comparées à celles des environnements marins actuels. Les pagures basaux
à carapaces cylindriques prédominaient dans les milieux récifaux du Jurassique supérieur, mais ont
été progressivement remplacés par des bernard-l’hermite à carapaces non cylindriques. Les pagures
les plus basaux révèlent un sillon branchial, mais la condensation branchiale vers un seul sillon (i.e.,
le cervical) est ici démontrée pour avoir eu lieu tôt dans leur histoire évolutive. Chez plusieurs genres
modernes, les restes d’un sillon branchial sont visibles, en combinaison avec plusieurs autres caractères
basaux. Pour cette raison, ceux-ci sont ici considérés comme les membres les plus primitifs parmi
les pagures actuels. Dans la présente étude, nous rétablissons la famille Probeebeidae, et érigeons la
nouvelle famille Paguropsidae n. fam, pour accueillir des bernard-l’hermite basaux actuels comme
Paguropsis Henderson, 1888, Eopaguropsis Fraaije, Krzemiński, Van Bakel, Krzemińska & Jagt, 2012
et Paguropsina Lemaitre, Rahayu & Komai, 2018. Sont également décrites une paire de pinces iso-
chèles de pagure du Jurassique inférieur (Pliensbachien) et deux paires de pinces isochèles de pagures
MOTS CLÉS du Jurassique moyen (Callovien), toutes recueillies récemment. Un nouveau spécimen de Schobertella
Condensation
branchiale, révèle certains traits morphologiques importants du bouclier qui n’avaient pas été observés auparavant.
isochélie, Le plus ancien exemple connu, à ce jour, de pinces clairement hétérochèles dans le registre fossile
hétérochélie, des bernard-l’hermite, provient des couches du Kimméridgien supérieur/Tithonien du Boulonnais
Probeebeidae,
Paguropsidae n. fam, (Nord-Ouest de la France). Pour la première fois, un schéma phylogénétique des Paguroidea marins,
famille nouvelle. basé sur des données morphologiques de carapaces, est présenté.
INTRODUCTION of evolutionary patterns and processes amongst paguroids.
The identification of morphological novelties during the
The classification of the superfamily Paguroidea Latreille, 1802, course of their Mesozoic evolution adds important data in
or hermit crabs, is by far the most convoluted amongst decapod unravelling their history.
crustaceans (McLaughlin et al. 2010) and fossil material has Following the landmark study by Van Bakel et al. (2008), over
only rarely been considered in this respect. As a result of 55 carapace-based Mesozoic paguroid taxa have been recorded
meticulous collecting from upper Mesozoic (Lower Jurassic- (Fraaije et al. 2019, 2020b) and assigned to eight families. Of
Upper Cretaceous) sedimentary rocks over the last twenty these, the Gastrodoridae Van Bakel, Fraaije, Jagt & Artal, 2008,
years, our insight into the diversity and evolutionary history of the Pilgrimchelidae Fraaije, 2014 and the Schobertellidae
hermit crabs has greatly increased, despite the fact that details Schweigert, Fraaije, Havlik & Nützel, 2013 comprise exclusively
of paguroid evolution remain elusive. However, observations Mesozoic representatives (Fraaije et al. 2019). With at least 18
on morphological changes in (functional) hard parts can species, in eight families, the Tithonian (Late Jurassic) paguroid
contribute in several ways to improving our understanding fauna from Ernstbrunn (Austria) is the most diverse extinct
2 GEODIVERSITAS • 2022 • 44 (1)
The evolution of hermit crabs on the basis of carapace morphology
C
B
A
FIG. 1. — Branchial condensation in basal paguroid genera: A, Platykotta Chablais, Feldmann & Schweitzer, 2011; B, Schobertella Schweigert, Fraaije, Havlik &
Nützel, 2013; C, Eopaguropsis Fraaije, Krzemiński, Van Bakel, Krzemińska & Jagt, 2012.
assemblage so far (Fraaije et al. 2019). Other notable Mesozoic 2. the presence of a long rostrum;
hermit crab faunules have been recorded from the Oxfordian 3. the possession of a well-ornamented carapace;
(Upper Jurassic) of southern Poland (Fraaije et al. 2012b, c, 2014a, 4. the presence of a mid-dorsal ridge on the shield;
b), the Kimmeridgian (Upper Jurassic) of southern Germany 5. the presence of a mesogastric process;
(Van Bakel et al. 2008; Fraaije 2014; Fraaije et al. 2017), the 6. the possession of a well-calcified posterior carapace;
Tithonian-Berriasian (Upper Jurassic-lowermost Cretaceous) of 7. the presence of a calcified, delineated cardiac region.
the Czech Republic (Fraaije et al. 2013c, 2020b), and the upper In view of the above, it is highly unfortunate that, to our
Albian (mid-Cretaceous) of northwest Spain (Fraaije et al. 2008, knowledge, there is not a single study of extant paguroids
2009, 2012a). in which details of carapace morphology are considered
Our novel palaeontological data provide important addi- (Fraaije et al. 2019). As a consequence, there is quite a lot
tions to molecular analyses and morphological descriptions of confusion in the literature regarding the nomenclature of
of internal organs published by biologists in order to unravel carapaces regions (and their delineation) in hermit crabs (see
the phylogeny and taxonomy of paguroids, which were aptly e.g., Morgan & Forest 1991). On the basis of new data from
referred to as the ‘paguroid puzzle’ by McLaughlin et al. (2010). the fossil record and, in particular, of the notion of ‘branchial
The molecular study by Tsang et al. (2011) and our own work condensation’, described here, carapace delineation can be
on fossil taxa (Fraaije et al. 2013b; Fraaije 2014; present paper) understood much more easily and used in interpretation of
both demonstrate extant hermit crabs to have originated from both extinct and extant paguroids effectively. It is our hope
symmetrically handed ancestors with cylindrical carapaces. that the data presented here will stimulate biologists to start
In order to gain a better insight into evolutionary processes using carapace morphology in order to achieve a more robust
amongst paguroids, we here present an overview of all cur- phylogeny for the superfamily Paguroidea.
rently available palaeontological data with regard to faunal
compositions and notable changes in the morphology of both
carapaces and first pereiopods through geological time. Impor- BRANCHIAL CONDENSATION
tant basal morphological characters of carapaces observed in
the fossil record are the following: Fraaije et al. (2012c) were the first to document an important
1. the presence of a branchial groove; trend seen in the carapace morphology of early (i.e., Late
GEODIVERSITAS • 2022 • 44 (1) 3
Fraaije R. H. B. et al.
F
A
E
D
B
C
FIG. 2. — Branchial condensation in representatives of the families Parapylochelidae (A, B), Gastrodoridae (C), Xylopaguridae (D, E) and Pylojacquesidae (F).
Jurassic) paguroids. This concerns the fusion of the V-shaped the basal family Annuntidiogenidae Fraaije, 2014 (which
branchial and subcircular cervical groove (as seen in, for instance, includes the genus Paguristes Dana, 1851) have relatively long
the genera Platykotta Chablais, Feldmann & Schweitzer, 2011, rostra as well (Fig. 4C).
Eogastrodorus Van Bakel, Fraaije, Jagt & Artal, 2008 and
Gastrodorus von Meyer, 1864, into a single, sinuous groove UNIFORM CARAPACE ORNAMENTATION
(cervical groove) in Eopaguropsis Fraaije, Krzemiński, Van When considering overall carapace morphology and, in
Bakel, Krzemińska & Jagt, 2012. This merger of two trans- particular, the presence of large, rimmed orbital cavities,
verse carapace grooves into one, the cervical groove, is here the Triassic genus Platykotta may be interpreted as the most
referred to as ‘branchial condensation’ (Fig. 1); it occurs in all ancient paguroid known. This, as well as the Early Jurassic
paguroid lineages. In some fossil and extant representatives, genera Schobertella Schweigert, Fraaije, Havlik & Nützel, 2013
remnants of the branchial groove and/or small parts of the and Eogastrodorus and the Late Jurassic to mid-Cretaceous
mesobranchial regions in between the cervical and branchial Gastrodorus, all have uniformly granular to tuberculate shields
grooves can still be observed (see Figs 1B; 2). Traces of this and posterior carapaces (see e.g., Krzemińska et al. 2020).
double transverse groove system have been observed, amongst Amongst extant paguroids only Labidochirus Benedict, 1892,
others, in the extant families Parapylochelidae Forest, 1987, Probeebei, Tisea Morgan & Forest, 1991 and Tylaspis have
Xylopaguridae Gašparič, Fraaije, Robin & De Angeli, 2016 completely ornamented carapaces. In none of the other hermit
and Pylojacquesidae McLaughlin & Lemaitre, 2001 (Fig. 2). crab families recognised currently has uniform ornamentation
been observed.
REDUCTION IN LENGTH OF (SPINOSE) ROSTRUM
One of the most basal paguroid genera known to date, Gas- CARAPACE MIDLINE CREST
trodorus von Meyer, 1864, has a very long, spinose rostrum, In many Mesozoic paguroid families, a midline crest on the
with a midline crest that continues posteriorly towards the carapace has been documented, for instance, in the Gastro-
cervical groove. Amongst paguroid lineages in Mesozoic reefal doridae (stratigraphical range: Bajocian [Middle Jurassic] to
settings, as inferred here, a rapid decrease in rostrum length Albian; see Van Bakel et al. 2008; Klompmaker et al. 2011;
has become apparent (Figs 2; 3). Krzemińska et al. 2016). As far as the family Parapylochelidae is
Amongst extant paguroids only Probeebei Boone, 1926 concerned, there are examples of such crests in Kimmeridgian
(Boone 1926a), Tylaspis Henderson, 1885 and Paguropsis (Fraaije 2014: fig. 9d) and Albian members (Fraaije et al. 2012a:
Henderson, 1888 have relatively long rostra with a midline fig. 5b), while there are also Kimmeridgian taxa assigned to the
crest. Apparently, a long spinose rostrum was advantageous families Pilgrimchelidae and Pylochelidae (Fraaije 2014: figs 7a;
only in certain deep-water habitats. Interestingly, but not 9e), as well as diogenids of Oxfordian (Fraaije et al. 2012c:
unexpectedly, of the remaining extant groups, members of fig. 2b, e), Tithonian (Fraaije et al. 2019: fig. 2c-f ) and Albian
4 GEODIVERSITAS • 2022 • 44 (1)
The evolution of hermit crabs on the basis of carapace morphology
C
B
A
FIG. 3. — Reduction of rostral length during the Late Jurassic in representatives of the families Gastrodoridae (A), Parapylochelidae (B) and Pylochelidae (C).
age (Fraaije et al. 2012a: fig. 5a). The oldest members of the CALCIFIED AND DELINEATED CARDIAC REGION
Paguridae known to date, of Tithonian age, also reveal a Gastrodorids have a well-calcified and well-delineated car-
midline crest on the shield (Fraaije et al. 2019: figs 4a-c; 5b). diac region, while this region is in part well delineated in
In modern hermit crabs, only Probeebei, Tylaspis and Paguropsis parapylochelids, probeebeids and paguropsids. Members
show a midline crest on the rostrum that continues, to a of all other hermit crab families lack this character.
greater or lesser extent, onto the shield.
TRENDS WITHIN PAGUROID FAMILIES
MESOGASTRIC ANTERIOR PROCESS Within the various families we have observed changes in a
In the Jurassic families Gastrodoridae, Schobertellidae, Dio- range of morphological features over geological time. For
genidae Ortmann, 1892, Paguropsidae n. fam (see below) instance, in annuntidiogenids, a widening of the frontal
and Parapylochelidae, a mesogastric anterior process is often shield has been observed in a number of species. In addi-
present (see e.g., Fraaije et al. 2019: figs 2g-i; 3a-e; 4d). In tion, the massetic region becomes bulkier and changes
modern paguroids, we have not been able to document any shape, from an elongated strip towards a more rectangular,
mesogastric anterior processes. globose part of the shield (Fig. 4).
Another trend is seen in the formation of, and a change
WELL-CALCIFIED POSTERIOR CARAPACE in shape in, an intragastric Y-linea within the Calcinidae
In general, members of the families Gastrodoridae, Fraaije, van Bakel & Jagt, 2017 (Fraaije et al. 2017: fig. 2,
Paguropsidae n. fam (see below), Parapylochelidae, Probeebeidae, 2020a). Unfortunately, the fossil record of paguroid cara-
Pylochelidae and Schobertellidae have relatively well-calcified paces of Paleogene and Neogene age is extremely scanty, with
posterior carapaces. Generally speaking, such are absent in all only a handful of specimens having been described to date.
other families. Needless to say, this hampers our understanding of paguroid
GEODIVERSITAS • 2022 • 44 (1) 5
Fraaije R. H. B. et al.
C
B
A
FIG. 4. — Changes in the massetic regions through time; from the Late Jurassic Annuntidiogenes jurassicus Fraaije, 2014 (A) to the mid-Cretaceous (late Albian)
Annuntidiogenes worfi Fraaije, van Bakel, Jagt, Klompmaker & Artal, 2009 (B) and extant Paguristes sp. (C).
evolution during the Cenozoic (e.g., Beschin et al. 2016; The first example of a hermit crab with a clear pair of
Fraaije et al. 2020a; Wallaard et al. 2020). heterochelous chelae (Fig. 7) is now contained in a private
collection; it has been collected from the upper Kimmerid-
PAGUROID ISOCHELY AND HETEROCHELY IN THE FOSSIL RECORD gian-lower Tithonian of the Atlantic Coast near Wimereux
To date, we are aware of only five examples of paired (iso- (Pas-de-Calais, northwest France). On the basis of a few
chelous) hermit crab chelae of Early and Middle Jurassic age, photographs of this particular specimen that we have
as follows: received from the collector, we have been able to document
– The holotype of Schobertella simonsenetlangi Schweigert, that the major right chela is about 1.5 to 2 times larger
Fraaije, Havlik & Nützel, 2013, and another specimen of the than the left one.
same species (Fig. 5) described herein (see below), both from There is a match, in geological time, between this presence
the upper Pliensbachian of southern Germany; of heterochelous chelae in a Late Jurassic paguroid and the first
– Orhomalus spinosus Schweitzer, Feldmann & Lazăr, 2009, occurrences of carapaces of members of the ‘asymmetrically
from the lower Callovian of Romania, and two newly collected handed’ Annuntidiogenidae, Diogenidae and Paguridae.
specimens from the middle Callovian of France (Fig. 6). These Interestingly, the majority of extant primitive genera (e.g.,
individuals all reveal isochelous or near-isochelous chelae; they Labidochirus, Paguropsis, Paguropsina Lemaitre, Rahayu &
were collected during field work along the high-speed railway Komai, 2018, Probeebei and Tisea ) also have equal or
line LGV SEA (Ligne à Grande Vitesse Sud Europe Atlan- subequal chelae. As a matter of fact, cheliped morphology
tique) by X. Valentin’s team, from the Gratte-Loup limestones in Labidochirus , Probeebei and Tisea resembles that of
near Chasseneuil-du-Poitou (Vienne, France). These levels are Cryptopagurus (see Schweigert et al. 2013: fig. 10a),
dated as middle Callovian (Coronatum ammonite Zone; see whereas that of Paguropsis and Paguropsina is close to that
Cariou 1980) and are situated near the historical quarries of in Schobertella (see Schweigert et al. 2013: figs 3-5; 9).
Grand Pont, Bonnillet and Lourdines (Mathieu 1968). These The chelae of Schobertella simonsenetlangi were described
specimens were associated with abundant bivalves, remains in detail by Schweigert et al. (2013). They are convex both
of marine crocodyliforms, plesiosaurs, sharks and bony fish, externally and internally and covered with about 20 irregular
as well as terrestrial plants. These two pairs of chelae will be longitudinal rows of variably sized, forwardly directed
the subject of a forthcoming taxonomic study. tubercles on either side. The fixed fingers and dactyli are
6 GEODIVERSITAS • 2022 • 44 (1)
The evolution of hermit crabs on the basis of carapace morphology FIG. 5. — Isochelous chelae of Schobertella simonsenetlangi Schweigert, Fraaije, Havlik & Nützel, 2013 (SMNS 70555, Staatliches Museum für Naturkunde Stuttgart, Germany, from the Lower Jurassic (upper Pliensbachian) Amaltheenton Formation of Iggingen near Aalen, southwestern Germany. Scale bars: 5 mm. A B FIG. 6. — Isochelous chelae of paguroids from the middle Callovian of Poitou, France: A, specimen UP/CGL.12.10 (UP = Université de Poitiers, CGL = Calcaires de Gratte-Loup). B, specimen UP/CGL.12.11. Photographs: Philippe Loubry. Scale bars: 10 mm. GEODIVERSITAS • 2022 • 44 (1) 7
Fraaije R. H. B. et al. FIG. 7. — The oldest example of heterochelous paguroid claws from the Upper Jurassic of the Wimereux area (Boulonnais, northwestern France); an individual preserved in situ in the internal mould of a Vetigastropoda (private collection of Mr Gilles Dron). The maximum diameter of the gastropod is about 7 cm. stout and compact with inwardly curved tips. The dentition, spine; a prominent central gastric process ending in a deep especially with extremely strong, elongate teeth close to incision at the base of the rostrum, a tapered and rounded the distal end of the fingers, points to a shell-cracking anterolateral border and tuberculate ornamentation. This function. A recently collected new specimen of Schobertella new specimen hints at a close relationship with the early simonsenetlangi reveals well-preserved details of the frontal paguropsid Eopaguropsis nidiaquilae Fraaije, Krzemiński, part of the shield (Fig. 8A): a triangular rostrum with Van Bakel, Krzemińska & Jagt, 2012, from the Oxfordian large subcircular orbital rim ending in blunt outer orbital of Poland (Fig. 8B). 8 GEODIVERSITAS • 2022 • 44 (1)
The evolution of hermit crabs on the basis of carapace morphology
A
B
FIG. 8. — Schobertella simonsenetlangi Schweigert, Fraaije, Havlik & Nützel, 2013 (SMNS 70555; see also Fig. 5), from the Lower Jurassic (upper Pliensbachian)
Amaltheenton Formation of Iggingen near Aalen, southwestern Germany: (A) showing new details of shield morphology, compared to Eopaguropsis nidiaquilae
Fraaije, Krzemiński, Van Bakel, Krzemińska & Jagt, 2012 (B) (see Fraaije et al. 2012c: fig. 2d). Scale bars: 10 mm.
PAGUROID FAUNAL When percentages for these three groups are plotted (see
COMPOSITION THROUGH TIME Fig. 9), a significant decline in cylindrical paguroids, coupled
with an increase in the numbers of non-cylindrical ones,
In order to document changes in paguroid faunal composition can be noted. In modern reefal settings, there are no basal
through time we have used three groups, namely the family paguroids; instead, a predominance of asymmetrically handed
Gastrodoridae, paguroids with ‘cylindrical carapace’ (i.e., all paguroids is apparent. The most basal taxa amongst extant
families depicted in Figure 12, to the left of the Gastrodori- paguroids, such as parapylochelids, pylochelids, paguropsids
dae) and other (‘non-cylindrical’) hermit crabs (i.e., those to and probeebeids, survived only in a few bathyal habitats and,
the right of the Gastrodoridae). in this way, are comparable to ‘Lazarus’ lobster taxa from
The oldest known paguroid faunule based on carapace remains is such settings, such as Neoglyphea inopinata Forest & de Saint
that from the Oxfordian of southern Poland (Van Bakel et al. 2008; Laurent, 1975, Laurentaeglyphea neocaledonica Forest, 2006
Fraaije et al. 2012b, c, 2014a; Krzemińska et al. 2016). This and polychelids (Polychelida).
comprises a single species of gastrodorid, three pilgrimchelids,
two pylochelids and a single paguropsid. RESURRECTION OF THE FAMILY PROBEEBEIDAE BOONE, 1926
The Kimmeridgian assemblage from Nusplingen (southern Th e monotypic genus Probeebei , and its type species,
Germany) has also been documented in detail in recent years P. mirabilis, were originally described as a primitive macruran
(Fraaije 2014) and comprises at least two gastrodorids, three by Boone (1926a). In the same year, Boone (1926b) erected
parapylochelids, two pilgrimchelids, three pylochelids, as well a new family, the Probeebeidae, to accommodate this
as one annuntidiogenid and one diogenid each. enigmatic crustacean. Much later, Wolff (1961) provided
Tithonian faunules from Ernstbrunn (Austria) are the most a detailed redescription of Probeebei and demonstrated
diverse extinct assemblage recorded to date (Fraaije et al. 2019), that it was a hermit crab, which he assigned to the family
with two gastrodorids, three pylochelids, two pilgrimchelids, Paguridae Latreille, 1802. The genus Probeebei was
two parapylochelids, one schobertellid, two annuntidiogenids, later transferred to the Parapaguridae Smith, 1882, by
four diogenids, one pagurid and two paguropsids. de Saint Laurent (1972). However, that author barely
From Tithonian-Berriasian strata in Moravia (Czech Republic), considered the carapace morphology of this genus, yet
Fraaije et al. (2013c, 2020a) have described an assemblage based her assignment to the Parapaguridae on features
that includes a single gastrodorid, two parapylochelids, one such as the presence of a labral spine, the absence of an
pilgrimchelid, two pylochelids, one diogenid, one paguropsid exopodal flagellum on the first maxilliped, the presence of
and two pagurids. undivided abdominal tergites and an unpaired left gonopore
So far, only a single mid-Cretaceous faunule is on record in females and the lack of a median constriction on the
from the upper Albian of northwest Spain (Fraaije et al. 2008, telson (compare Lemaitre 1998). None of these characters
2009, 2012a), comprising one gastrodorid, one parapylochelid, was used in the descriptions of Probeebei and Tylaspis by
one pylochelid and two annuntidiogenids. de Saint Laurent (1972) and are not found either in any
GEODIVERSITAS • 2022 • 44 (1) 9Fraaije R. H. B. et al.
Extant reefs
Albian Koskobilo
Tithonian Ernstbrunn
Kimmeridgian Nusplingen
Oxfordian Poland
FIG. 9. — Changes in taxonomic composition amongst paguroid assemblages within reefal settings during the Late Jurassic (Oxfordian, Kimmeridgian and Titho-
nian) and mid-Cretaceous (Albian) compared to modern faunas. Grey, Gastrodoridae; orange, cylindrical carapace; blue, non-cylindrical carapace.
10 GEODIVERSITAS • 2022 • 44 (1)The evolution of hermit crabs on the basis of carapace morphology
E
D
C
B
A
FIG. 10. — Transition from the Gastrodoridae to the Probeebeidae, featuring branchial condensation, reduction of rostrum, reduction of ornament on shield, inflation of
posterior carapace, reduction of delineation of cardiac and reduction of midline carapace crest: A, Gastrodorus; B, Probeebei; C, Labidochirus; D, Tylaspis; E, Tisea.
of the more recent accounts on parapagurid taxa (see e.g., Family PROBEEBEIDAE Boone, 1926, emended herein
Lemaitre 2013, 2014). An unpaired left gonopore is also
a common trait in the Paguridae (e.g., Lemaitre 2003). Probeebeidae Boone, 1926b: 73.
The monotypic genera Probeebei and Tylaspis were TYPE GENUS. — Probeebei (Boone, 1926) by monotypy.
restudied and redescribed in detail by Lemaitre (1998),
who noted that de Saint Laurent (1972) had erroneously OTHER GENERA INCLUDED. — Labidochirus Benedict, 1892, Tisea
Morgan & Forest, 1991 and Tylaspis Henderson, 1885.
indicated that she had examined specimens of Tylaspis
anomala Henderson, 1885 from the Indian Ocean. In DIAGNOSIS. — Shield (excluding rostrum) strongly convex, well
actual fact, this material had originated from the Pacific calcified, width equal to length or width exceeding length, with
Ocean. Lemaitre (1998) also stated that several important distinct bulges (i.e., keraial and massetic regions) laterally, strong
to weaker spinose ornament. Rostrum well developed, exceeding
morphological features of T. anomala had been inaccurately lateral projections. Cervical and branchial grooves subparallel, en-
or insufficiently recorded in previous studies and found compassing small branchial regions. Posterolateral margins spinose.
additional discrepancies or inaccuracies in published accounts Posterior carapace well calcified, broadly inflated, with dense spinose
of Probeebei and Tylaspis. He also observed a similar mode of ornamentation. Well-developed cardiac grooves encompassing
life and identical morphological characters of the carapace cardiac region.
(different from those of other parapagurids) for these two REMARKS. — The bulges on the shield of extant probeebeids (Fig. 10)
crab-like genera and assumed these two paguroids to be are quite similar to those of the most basal paguropsid, Eopaguropsis
advanced forms. Our data show these two taxa retain almost nidiaquilae (see Fraaije et al. 2012c: figs 2c; 3), which also has a
all of the ‘remnant’ basal carapace characteristics mentioned spinose ornamentation on the gastric and lateral parts of the shield.
above. For this reason, we consider them to be the most
basal ones amongst extant paguroids. In fact, together with ERECTION OF A NEW FAMILY, PAGUROPSIDAE N. FAM.
Parapylocheles Alcock, 1901, they are amongst the most Paguropsis Henderson, 1888 has remained a monotypic genus
primitivex of modern-day hermit crabs. ever since its original description. In spite of the uniqueness
GEODIVERSITAS • 2022 • 44 (1) 11Fraaije R. H. B. et al.
C
B
A
FIG. 11. — Paguropsids (Paguropsidae n. fam) through time, from the Oxfordian Eopaguropsis Fraaije, Krzemiński, Van Bakel. Krzemińska & Jagt, 2012 (A), via the
Tithonian Eopaguropsis (B) to extant Paguropsis Lemaitre, Rahayu & Komai, 2018 (C).
of this genus amongst the Paguroidea, the taxonomy and on cheliped coxae has also been documented in species of
morphology of Paguropsis typica have not been studied in any Paguristes (Lemaitre et al. 2018). Alcock (1905) had already
detail that would meet the requirements of modern paguroid indicated that Paguropsis was closely related to Paguristes.
studies (Lemaitre et al. 2018). Lemaitre et al. (2018) noted Species of Paguropsis and Paguropsina are all symbionts of
that Paguropsis was not monotypic any longer, and that in acontiate anemones (Actiniaria) (Lemaitre et al. 2018). This
addition to P. typica, there was an additional genus and six type of association and habitat can potentially be postulated
species. The striking symmetry of the carapace, pereiopods for Mesozoic counterparts (Eopaguropsis spp.) as well.
(including chelipeds), uropods and telson, are unique
amongst the so-called “asymmetrically handed” paguroids.
In illustrations provided by Lemaitre et al. (2018: figs 2a-d Family PAGUROPSIDAE n. fam.
and 18d, in particular), important, yet hitherto unobserved
and/or unrecorded paguroid carapace features are shown urn:lsid:zoobank.org:act:88399E62-3864-4B66-8D89-FC32C562F5A9
for Paguropsis and Paguropsina. These features are: deep slits
along the postfrontal ridges centrally posterior to the orbital TYPE GENUS. — Paguropsis Henderson, 1888.
cavity and centrally directed incisions near the cervical groove
OTHER GENERA INCLUDED. — Eopaguropsis Fraaije, Krzemiński,
between the posterior margin and branchial area in the central Van Bakel. Krzemińska & Jagt, 2012 and Paguropsina Lemaitre,
part of the narrow lateral carapace lobe. These features are also Rahayu & Komai, 2018 (Fig. 11).
seen in an Eocene paguroid from Italy (Beschin et al. 2021).
A decalcified longitudinal fissure on the ventral face of the ETYMOLOGY. — The name is derived from the type genus.
coxae of the chelipeds occurs in all species of Paguropsis and DIAGNOSIS. — Prominent subtriangular rostrum, considerably
Paguropsina. A similar, and presumably homologous, fissure exceeding lateral projections. Cervical and branchial grooves subparallel,
12 GEODIVERSITAS • 2022 • 44 (1)The evolution of hermit crabs on the basis of carapace morphology
Pylojacquesidae
Parapaguridae
Xylopaguridae
Probeebeidae
Diogenidae
Calcinidae
Paguridae
Cenozoic
6
Cretaceous
Paguropsidae n. fam.
Pylochelidae
Pilgrimchelidae
Parapylochelidae
Schobertellidae
Gastrodoridae
Annuntidiogenidae
Jurassic
7
4
3 5
Triassic
2 1
FIG. 12. — Intuitive phylogenetic tree based on carapace morphologies as outlined in the present study and on data from McLaughlin et al. (2007) and Fraaije et al. (2019).
Important synapomorphies include the following: 1, cylindrical carapace; 2, cervical and branchial grooves widely separated; 3, arrow-shaped gastric region;
4, cervical groove not extending to lateral border; 5, intragastric grooves; 6, Y-linea; 7, spinose posterolateral borders.
encompassing small branchial regions (= lateral lobes) or forming broad, novel niches through time, shifts in carapace and cheliped
sinuous groove. Shield well calcified, subtriangular or subrectangular; morphologies occurred. A better-adapted metabolism and
dorsal surface slightly vaulted with incomplete midline crest; well-
delineated massetic region covered with numerous setal pits; posterior
changes of food source also had an impact on gill structure
carapace less calcified, delineated cardiac region; uropods and telson and internal musculature and consequently led to internal
symmetrical. Chelipeds subequal, similar in armature and setation. rearrangement (e.g. Van Bakel et al. 2012). Additional mate-
rial and more in-depth studies are needed to document the
increased diversity in morphology, physiology and behaviour
PHYLOGENY OF THE PAGUROIDEA within the Paguroidea during deep time.
BASED ON CARAPACE MORPHOLOGY
Based on the data presented above and the stratigraphical ranges Acknowledgements
of extinct paguroid taxa based on carapaces, a phylogeny of Our late colleague Patsy McLaughlin opened our eyes when,
all marine paguroids is here proposed (Fig. 12). around 2007, we embarked on our studies of extinct her-
Overall, it can be concluded that branchial condensa- mit crabs, by hinting at the close relationships of our Late
tion within the Paguroidea appeared early in the Jurassic; Jurassic material with the genera Paguropsis and Tylaspis
it has also been recognised (and used for taxonomic pur- and members of the family Pylochelidae; her assistance
poses) in a wide array of Mesozoic brachyurans (Van Bakel was instrumental in producing our 2008 paper. Dr Charles
et al. comm. pers.). Recognition and documentation of this Fransen (Naturalis Biodiversity Center, Leiden, the Nether-
phenomenon facilitate our understanding of paguroid car- lands) supplied several items of literature for which we are
apace delineation. Together with other carapace characters, grateful. G. G. and X. V. are appreciative to the municipality
cheliped size and trends through time, as outlined above, the of Chasseneuil-du-Poitou (Mr C. Eidelstein) and to Mrs E.
fossil data have contributed to the family tree of marine her- Blandin, P. Ferchaud and J. M. Terrasson for field assistance.
mit crabs presented here. The occurrence of paired isochelous Their work is supported in part by the Lisea Fondation
chelae and symmetrical calcified tergites in extinct paguroids Biodiversité (research grant 97 to Palaios association with
(Fraaije et al. 2012d, 2013c, 2014b) provides strong support the project “Past of Biodiversity from Vienne”). Hemke
for the symmetrical ancestry of the Paguroidea. Dekkers, Shjalina Lentink, Mila van Loon, Lars Rongen,
Major Mesozoic evolutionary and radiation events amongst all students from SintLucas Vakschool voor creatief talent,
marine phyto- and zooplanktonic microbiota led, both directly Eindhoven, assisted in finishing all figures. Finally, com-
and indirectly, to a rise of new ecological niches and, concom- ments made by Rok Gašparič (Kamnik/Ljubljana, Slovenia)
itantly, development of various marine benthic and nektonic and an anonymous reviewer on an earlier version of the
groups (Fraaije et al. 2018). Linked to the occupation of these typescript are much appreciated.
GEODIVERSITAS • 2022 • 44 (1) 13Fraaije R. H. B. et al.
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